Manx Shearwaters in the Bay of Biscay

Prior to forming the egg, female Manx Shearwaters from south Irish Sea colonies spend a fortnight or so exploiting the rich sardine fishery in the Bay of Biscay; the males remain behind, in nightly occupation of the burrows.     

Temporal patterns in the acoustic signals of beaked whales at Cross Seamount

Seamounts may influence the distribution of marine mammals through a combination of increased ocean mixing, enhanced local productivity and greater prey availability. To study the effects of seamounts on the presence and acoustic behaviour of cetaceans, we deployed a high-frequency acoustic recording package on the summit of Cross Seamount during April through October 2005. The most frequently detected cetacean vocalizations were echolocation sounds similar to those produced by ziphiid and mesoplodont beaked whales together with buzz-type signals consistent with prey-capture attempts. Beaked whale signals occurred almost entirely at night throughout the six-month deployment. Measurements of prey presence with a Simrad EK-60 fisheries acoustics echo sounder indicate that Cross Seamount may enhance local productivity in near-surface waters. Concentrations of micronekton were aggregated over the seamount in near-surface waters at night, and dense concentrations of nekton were detected across the surface of the summit. Our results suggest that seamounts may provide enhanced foraging opportunities for beaked whales during the night through a combination of increased productivity, vertical migrations by micronekton and local retention of prey. Furthermore, the summit of the seamount may act as a barrier against which whales concentrate prey.     

Growth of young-of-the-year mackerel in the Bay of Biscay

The first growth season of young-of-the-year (0+ year) mackerel Scomber scombrus , sampled in the Bay of Biscay, was parameterized to determine growth patterns. Daily increments were identified on sagittae otoliths, for calculation of age and growth of 92 larvae and 54 juveniles over the range 3·6–215·0 mm standard length ( L _S). A Gompertz curve was fitted to the length-at-age data. At the end of the first year of growth L _S was 194·2 mm, with a maximum growth increment of c . 2 mm day⁻¹, observed 62 days after hatching. Backcalculated growth increments for mackerel juveniles, during their larval stage, were higher than those observed for sampled larvae; only 10·9% of sampled larvae were estimated to survive. Growth for north-eastern Atlantic mackerel was slower than that published for north-western Atlantic mackerel. Backcalculated hatching dates for mackerel were consistent with the typical temporal distribution of mackerel spawning in the Bay of Biscay.     

No shallow talk: Cryptic strategy in the vocal communication of Blainville's beaked whales

Communicating animals must balance fitness benefits against the costs of signaling, such as increased predation risk. Cetaceans communicate mainly with sound and near‐surface vocalizations can place signalers at risk from shallow‐diving top‐predators with acute hearing such as killer whales. Beaked whales are deep divers living in small cohesive groups with little social defense from predation. Little if anything is known about their acoustic communication. Here, eight Blainville's beaked whales were studied with suction cup attached DTags to provide the first report on social communication as a function of diving behavior for any of the 21 ziphiid species. Tagged whales produced two previously unrecorded signals with apparent communicative functions: (1) fast series of ultrasonic frequency modulated clicks (rasps) were recorded from six individuals, and (2) harmonically rich short whistles with a mean fundamental frequency of 12 kHz were recorded from one whale at up to 900 m depth, the deepest whistles recorded from a marine mammal. Blainville's were silent 80% of the time, whenever shallower than 170 m depth and during the prolonged (19 min) silent ascents from vocal dives. This behavior limits the ability of shallow‐diving predators to track Blainville's acoustically and may provide a striking example of the evolutionary influence of the risk of predation on animal communication.     

Rostral densification in beaked whales: Diverse processes for a similar pattern

As compared to other odontocetes (toothed whales), the rostrum of beaked whales (family Ziphiidae) often displays extensive changes in the shape, thickness, and density of its constituent bones. Previous morphological observations suggested that these modifications appeared in parallel in different ziphiid lineages. However, very few data were available on the compactness and histology of these rostral bones, which precluded the study of the processes at work for the development of such structures, as well as the interpretation of their functional implications. In this work we review the bibliographic data on the anatomy of the ziphiid rostrum and we add new observations on adults of several extinct and extant taxa. These observations are based on CT scans and transverse histological sections. Our results confirm that different bones (vomer, mesethmoid, premaxilla, maxilla) are involved in the various morphologies displayed by ziphiid rostra. Strong density contrasts are detected between bones and/or inside the bones themselves; for example, parts of the rostrum reach densities in the range of Neoceti ear bones, which are among the densest bones known hitherto. Furthermore, the histology of the pachyostotic and osteosclerotic bones proves to change from one taxon to the other; the degree of Haversian remodeling varies strongly between species: it can be absent (e.g. Aporotus recurvirostris), partial (e.g. aff. Ziphirostrum), or complete (e.g., Mesoplodon densirostris). The atypical secondary osteons known to be responsible for bone hypermineralization in the rostrum of M. densirostris occurred also in Choneziphius sp. Confronted with a phylogenetic framework, these anatomical and histological observations indicate that the acquisition of compact (osteosclerotic) and/or swollen (pachyostotic) bone is the result of a broad convergence between taxa, in response to common selective pressures. The functional dimension of this question is discussed with respect to what is known about extant ziphiid ecology.     

Gastrointestinal helminths of Cuvier's beaked whales, Ziphius cavirostris, from the western Mediterranean

We examined the gastrointestinal helminth fauna of 2 Cuvier's beaked whales, Ziphius cavirostris, stranded on the Spanish Mediterranean coast. Information regarding intestinal parasites of this species is provided for the first time. Six helminth taxa were identified. Thirty type II larvae of the nematode Anisakis sp. were found in the stomach and the intestine of both hosts; 2 type I larvae of Anisakis sp. were found in the intestine of 1 host. One juvenile of the acanthocephalan Bolbosoma vasculosum was found in the intestine; the metacestode Scolex pleuronectis was found mainly in the terminal colon and the anal crypts of both hosts; adult cestodes of Tetrabothrius sp., which may represent a new species, were collected from the duodenum of 1 host. Composition of the intestinal parasitic community is similar to that of other oceanic cetaceans, which mostly include species of Bolbosoma and tetrabothriids (Cestoda).     

A comprehensive and validated molecular taxonomy of beaked whales, family Ziphiidae

DNA sequences from orthologous loci can provide universal characters for taxonomic identification. Molecular taxonomy is of particular value for groups in which distinctive morphological features are difficult to observe or compare. To assist in species identification for the little known family Ziphiidae (beaked whales), we compiled a reference database of mitochondrial DNA (mtDNA) control region (437 bp) and cytochrome b (384 bp) sequences for all 21 described species in this group. This mtDNA database is complemented by a nuclear database of actin intron sequences (925 bp) for 17 of the 21 species. All reference sequences were derived from specimens validated by diagnostic skeletal material or other documentation, and included four holotypes. Phylogenetic analyses of mtDNA sequences confirmed the genetic distinctiveness of all beaked whale species currently recognized. Both mitochondrial loci were well suited for species identification, with reference sequences for all known ziphiids forming robust species-specific clades in phylogenetic reconstructions. The majority of species were also distinguished by nuclear alleles. Phylogenetic comparison of sequence data from "test" specimens to these reference databases resulted in three major taxonomic discoveries involving animals previously misclassified from morphology. Based on our experience with this family and the order Cetacea as a whole, we suggest that a molecular taxonomy should consider the following components: comprehensiveness, validation, locus sensitivity, genetic distinctiveness and exclusivity, concordance, and universal accessibility and curation.     

中国大陆海域沿岸搁浅和误捕的喙鲸类调查

从2009 年1 月至2010 年2 月,我们对中国大陆海域沿岸搁浅和误捕的喙鲸类头骨标本进行了测量和鉴
定。目前中国大陆海域搁浅和误捕的喙鲸类有5 种,分别为柯氏喙鲸、贝氏喙鲸、朗氏喙鲸、柏氏中喙鲸和小
中喙鲸,其中小中喙鲸为中国海域(包括台湾)的首次报道,也是在西北太平洋区的首次记录。小中喙鲸的主
要形态鉴别特征为有1 对纺锤形牙齿着生在下颌骨联合后面。经对已报道过的大陆海域搁浅和误捕的3 个雌性
银杏齿中喙鲸头骨标本和台湾省搁浅银杏齿中喙鲸的颅骨、鼻骨、下颌骨以及牙齿特征对比重新鉴定,订正大
陆的3 个样本为雌性柏氏中喙鲸。本文报道了大陆沿岸海域搁浅和误捕的喙鲸类最新调查研究结果。     

Energy density of anchovy Engraulis encrasicolus in the Bay of Biscay

The energy density ( E _D) of anchovy Engraulis encrasicolus in the Bay of Biscay was determined by direct calorimetry and its evolution with size, age and season was investigated. The water content and energy density varied seasonally following opposite trends. The E _D g⁻¹ of wet mass ( M _W) was highest at the end of the feeding season (autumn: c . 8 kJ g⁻¹ M _W) and lowest in late winter ( c . 6 kJ g⁻¹ M _W). In winter, the fish lost mass, which was partially replaced by water, and the energy density decreased. These variations in water content and organic matter content may have implications on the buoyancy of the fish. The water content was the major driver of the energy density variations for a M _W basis. A significant linear relationship was established between E _D g⁻¹ ( y ) and the per cent dry mass ( M _D; x ): y =−4·937 + 0·411 x . In the light of the current literature, this relationship seemed to be not only species specific but also ecosystem specific. Calibration and validation of fish bioenergetics models require energy content measurements on fish samples collected at sea. The present study provides a first reference for the energetics of E. encrasicolus in the Bay of Biscay.     

Molecular genetic identification of southern hemisphere beaked whales (Cetacea: Ziphiidae)

To assist in the species-level identification of stranded and hunted beaked whales, we compiled a database of 'reference' sequences from the mitochondrial DNA control region for 15 of the 20 described ziphiid species. Reference samples for eight species were obtained from stranded animals in New Zealand and South Australia. Sequences for a further seven species were obtained from a previously published report. This database was used to identify 20 'test' samples obtained from incompletely documented strandings around New Zealand. Analyses showed that four of these 'test' specimens (20%) had initially been misidentified. These included two animals of particular interest: (i) a Blainville's beaked whale ( Mesoplodon densirostris) , the first record of this species in New Zealand waters; and, (ii) a juvenile Andrews' beaked whale ( Mesoplodon bowdoini ), a species known from just over 20 strandings worldwide. A published sequence from a beaked whale product purchased in the Republic of Korea was identified as a Cuvier's beaked whale ( Ziphius cavirostris ). Levels of intra- and interspecific variation were compared to determine the potential for misidentification when the database or taxonomy is incomplete. Intraspecific variation was generally <2%, and interspecific divergence was generally >4.7%. Exceptions were within-species variation in Hyperoodon planifrons , southern bottlenosed whale (4.12%), which exceeded the variation between the two species of Berardius (3.78%), and variation between the two specimens assigned to M. hectori , Hector's beaked whale (7.14%). The latter case appears to be an error in species identification, and could represent the discovery of a new species of beaked whale.     

First direct measurements of behavioural responses by Cuvier's beaked whales to mid-frequency active sonar

Most marine mammal­ strandings coincident with naval sonar exercises have involved Cuvier''s beaked whales (Ziphius cavirostris). We recorded animal movement and acoustic data on two tagged Ziphius and obtained the first direct measurements of behavioural responses of this species to mid-frequency active (MFA) sonar signals. Each recording included a 30-min playback (one 1.6-s simulated MFA sonar signal repeated every 25 s); one whale was also incidentally exposed to MFA sonar from distant naval exercises. Whales responded strongly to playbacks at low received levels (RLs; 89–127 dB re 1 µPa): after ceasing normal fluking and echolocation, they swam rapidly, silently away, extending both dive duration and subsequent non-foraging interval. Distant sonar exercises (78–106 dB re 1 µPa) did not elicit such responses, suggesting that context may moderate reactions. The observed responses to playback occurred at RLs well below current regulatory thresholds; equivalent responses to operational sonars could elevate stranding risk and reduce foraging efficiency.     

Adult male Cuvier's beaked whales (Ziphius cavirostris) engage in prolonged bouts of synchronous diving

Studies of the social behavior of Cuvier's beaked whales (Ziphius cavirostris) are challenging because of their deep-water habitat usually far from shore and the limited time they spend at the surface. The sociality of these deepest diving mammals is of interest, however, especially for our understanding of how social systems evolve in extreme habitats. High levels of scarring suggest that males compete agonistically for access to females and so we predicted that associations among adult males would be unstable due to competitive exclusion. We tested this prediction by evaluating the diving behavior of animals within social groups off Cape Hatteras, North Carolina, considering diving synchrony a proxy for group membership. Using data from satellite-linked depth-recording tags, we found that adult male–male pairs showed extended periods of synchrony in diving behavior, while all pairs that included an adult male with an individual of another age and/or sex dove synchronously for less than a day. We assessed three hypotheses to explain these surprising results: sexual segregation; extended bouts of male–male competition; and the presence of male alliances. Finally, we considered testable predictions to distinguish among these explanations.     

Occurrence and social structure of Baird's beaked whales,Berardius bairdii,in the Commander Islands,Russia

The social structure of Baird's beaked whales is completely unstudied, and it is unknown if either females or males form long‐term social associations or occur in stable groups. In this paper we summarize our observations of individually identified animals over the span of 6 yr to provide insight on their long‐term social structure. We have identified 122 whales, with 28 of them encountered three times or more and thus included in the analysis of social structure. We found that the whales exhibited nonrandom patterns of social associations with some individuals preferentially associating with each other. Whales with more scarred skin had higher maximum association coefficients, which indicates that older animals and/or males were more inclined to form stable associations. Cluster analysis with a modularity test for gregariousness divided the whales into four clusters. Whales from the same clusters did not always occur together, but some individuals retained stable associations over several years. The strength of social relationships decayed over periods of months, with between‐year relationships showing little deviation from what would be expected if association was random. Generally these findings do not correspond to a stable society with fixed groups but instead suggest a fission‐fusion society with some stable alliances.     

The extracranial arterial system in the heads of beaked whales,with implications on diving physiology and pathogenesis

Beaked whales are medium‐sized toothed whales that inhabit depths beyond the continental shelf; thus beaked whale strandings are relatively infrequent compared to those of other cetaceans. Beaked whales have been catapulted into the spotlight by their tendency to strand in association with naval sonar deployment. Studies have shown the presence of gas and fat emboli within the tissues and analysis of gas emboli is suggestive of nitrogen as the primary component. These findings are consistent with human decompression sickness (DCS) previously not thought possible in cetaceans. Because, tissue loading with nitrogen gas is paramount for the manifestation of DCS and nitrogen loading depends largely on the vascular perfusion of the tissues, we examined the anatomy of the extracranial arterial system using stranded carcasses of 16 beaked whales from five different species. Anatomic regions containing lipid and/or air spaces were prioritized as potential locations of nitrogen gas absorption due to the known solubility of nitrogen in adipose tissue and the nitrogen content of air, respectively. Attention was focused on the acoustic fat bodies and accessory sinus system on the ventral head. We found much of the arterial system of the head to contain arteries homologous to those found in domestic mammals. Robust arterial associations with lipid depots and air spaces occurred within the acoustic fat bodies of the lower jaw and pterygoid air sacs of the ventral head, respectively. Both regions contained extensive trabecular geometry with small arteries investing the trabeculae. Our findings suggest the presence of considerable surface area between the arterial system, and the intramandibular fat bodies and pterygoid air sacs. Our observations may provide support for the hypothesis that these structures play an important role in the exchange of nitrogen gas during diving. J. Morphol. 277:5–33, 2016. © 2015 Wiley Periodicals, Inc.     

Plankton distribution across a slope current-induced front in the southern Bay of Biscay

Relatively warm (12.50–12.75°C) and high-salinity[<35.640 practical salinity units (PSU)] water flowing eastwardwas detected at the shelf-break during a cruise carried Outin the southern Bay of Biscay in Spring 1987. The slope currentinduced the formation of a convergent front separating well-mixedoceanic waters from haline-stratified coastal waters. Very highconcentrations of dissolved oxygen (295 µmol kg^–1)and chlorophyll a(>4.5 mg m^–3) were found at the outeredge of the frontal boundary. Small autotrophic flagellatesdominated the phytoplankton community. Primary production peakedat the boundary region. Estimated phytoplankton growth ratesindicated that active growth was taking place, with lower turnovertimes integrated over the water column at the frontal station(2.5–5 days) than at coastal (1.5–2.8 days) or oceanic(1.5–3.5 days) stations. The lowest doubling times (1–2days) were calculated for surface frontal populations. Accumulationof zooplankton was also observed associated with the convergentphysical structure, although this relationship was less markedthan for phytoplankton. Copepods, mainly Paracalanus parvus,Acartia clausi and Oithona helgolandica, formed the bulk ofthe mesozooplankton biomass. Compatibility between the sizeof phytoplankton cells and copepod size spectra indicate highfood availability for these animals, particularly in the vicinityof the front. The distribution of fish eggs and fish larvaewas also coupled with the slope current-induced front. Sardinelarvae were more abundant at the coastal side of the front,whereas larval stages of blue whiting reached the highest densitiesat off-shelf stations. Larvae of lamellibranch molluscs andbryozoa were restricted to nearshore waters, as the frontalboundary prevented larval dispersion to the open ocean. Theresults presented in this paper suggest that the Iberian slopecurrent and its associated shelf-break frontal structure werecrucial in controlling phytoplankton primary production, activityof grazers, distribution of larvae of fishes and benthic invertebrates,and ultimately in determining the structure of the pelagic foodweb in the southern Bay of Biscay during the seasonal periodof vertical mixing.     

The extracranial venous system in the heads of beaked whales,with implications on diving physiology and pathogenesis

Beaked whales are a poorly known but diverse group of whales that have received considerable attention due to strandings that have been temporally and spatially associated with naval sonar deployment. Postmortem studies on stranded carcasses have revealed lesions consistent with decompression sickness, including intravascular gas and fat emboli. These findings have been supported by analyses of intravascular gas emboli showing composition dominated by nitrogen gas. To increase our understanding of the pathophysiology of nitrogen bubble formation and intravascular embolization, we examined the gross and microscopic anatomy of the venous system in the head of beaked whales. Since the potential sources of intravascular fat and gas emboli were of greatest interest, focus was placed on the acoustic fat bodies and pneumatic accessory sinus system. Herein, we describe intimate arteriovenous associations with specialized adipose depots and air sinuses in beaked whales. These vascular structures comprise an extensive network of thin‐walled vessels with a large surface area, which is likely to facilitate exchange of nitrogen gas and may, therefore, form anatomic regions that may be important in physiological management of diving gases. These structures may also be vulnerable to pathologic introduction of emboli into the vascular system. Expansive, thin‐walled venous lakes are found within the pterygoid region, which suggest the potential for nitrogen exchange as well as for compensation of middle‐ear pressures during descent on a dive. These findings warrant further research into the structure and function of this morphology as it relates to normal and pathologic physiology. J. Morphol. 277:34–64, 2016. © 2015 Wiley Periodicals, Inc.     

Following a foraging fish-finder: diel habitat use of Blainville's beaked whales revealed by echolocation

Simultaneous high resolution sampling of predator behavior and habitat characteristics is often difficult to achieve despite its importance in understanding the foraging decisions and habitat use of predators. Here we tap into the biosonar system of Blainville''s beaked whales, Mesoplodon densirostris, using sound and orientation recording tags to uncover prey-finding cues available to echolocating predators in the deep-sea. Echolocation sounds indicate where whales search and encounter prey, as well as the altitude of whales above the sea-floor and the density of organisms around them, providing a link between foraging activity and the bio-physical environment. Tagged whales (n = 9) hunted exclusively at depth, investing most of their search time either in the lower part of the deep scattering layer (DSL) or near the sea-floor with little diel change. At least 43% (420/974) of recorded prey-capture attempts were performed within the benthic boundary layer despite a wide range of dive depths, and many dives included both meso- and bentho-pelagic foraging. Blainville''s beaked whales only initiate searching when already deep in the descent and encounter prey suitable for capture within 2 min of the start of echolocation, suggesting that these whales are accessing prey in reliable vertical strata. Moreover, these prey resources are sufficiently dense to feed the animals in what is effectively four hours of hunting per day enabling a strategy in which long dives to exploit numerous deep-prey with low nutritional value require protracted recovery periods (average 1.5 h) between dives. This apparent searching efficiency maybe aided by inhabiting steep undersea slopes with access to both the DSL and the sea-floor over small spatial scales. Aggregations of prey in these biotopes are located using biosonar-derived landmarks and represent stable and abundant resources for Blainville''s beaked whales in the otherwise food-limited deep-ocean.     

The distribution of scleractinian corals in the Bay of Biscay, NE Atlantic

Studies on the distribution of scleractinian corals in the Bay of Biscay were relatively numerous during the nineteenth and the first half of the twentieth century. Yet, recent reports are scattered, sometimes unpublished, and therefore knowledge about the current coral occurrence in the area is limited. This study aims at compiling the available historical and more recently collected information on the occurrence of scleractinian corals in the Bay of Biscay. Data from two recent cruises are included and compared with previous explored coral sites from as early as 1830 up to 1995. A database of 347 records including 34 described species of scleractinians highlights that the slope of the Bay of Biscay is an important habitat for scleractinians. This could be expected due to the high topographic relief providing the necessary hard substrate and accelerated bottom current flow that corals require. Further exploration of the occurrence and ecology of corals in the area is recommended to support the conservation of cold-water coral reefs along the European margin. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

利用mtDNA序列及形态特征重新鉴定福建长乐搁浅喙鲸

<正>中国大陆沿岸属大陆棚水域,常见的齿鲸物种以近岸型的海豚科(Delphinidae)、小抹香鲸科(Kogiidae)及鼠海豚科(Phocaenidae)动物为主(王丕烈,2012),而喙鲸科(Ziphiidae)发现纪录相对较少,近年来仅由王丕烈等(2011)针对喙鲸科的中喙鲸属(Mesoplodon)标本进行检视校正,认为中国大陆沿岸存在柏氏中喙鲸